Leadership in Wolf, Canis lupus, Packs

Introduction

Leadership in Wolf packs has been addressed only a few times. Mech (1970:73)
defined leadership among Wolves as "... the behavior of one Wolf that obviously
controls, governs, or directs the behavior of several others" such as when Wolves
decide on direction of travel, when to rest or travel, and whether to chase
prey. Peterson (1977) and Haber (1977) adopted similar definitions.

However, few opportunities have existed for studying leadership in wild Wolf
packs because of the elusive nature of Wolves. Only Murie (1944), Clark (1971),
Haber (1977), and Mech (1988, 1995a, 1999) have studied behavior in free-ranging
Wolf packs, and no one has examined the leadership concept critically or quantitatively.
This article attempts to do so, based primarily on my 13 summers of observation
of a free-ranging Wolf pack.

Because Wolf packs are basically families (Murie 1944; Mech 1970), or at
least almost always include a breeding pair (Mech and Nelson 1990; Mech et
al. 1998), it is only natural that some member of this pair would be the pack
leader (Mech 1970). This is because most members of the pack would be the
offspring of the breeding pair and would tend to follow their parents' initiatives.
Sometimes, a post-reproductive animal remains with the pack (Mech 1995a).
Conceivably, such an individual, being older and more experienced, would lead
in some activities. On the other hand, deposed breeders usually become subordinate,
and tend not to take initiatives involving the group (Zimen 1976). A post-reproductive
female on Ellesmere was subordinate to the breeding pair (Mech 1999).

Therefore, determining which member(s) leads the pack in a given activity
would usually involve determining whether it is the breeding male or breeding
female. Murie (1944) identified a male "lord and master" of a pack to which
all four other adults submitted. Murie concluded that this animal was not
mated to any of the females, although there was no way he could have known
(Haber 1977). One of the other adult males tended to lead the chases of Caribou,
Rangifer tarandus, during Murie's study.

On Isle Royale, Mech (1966) observed from the air that in a large pack one
member stood out as leading the pack, taking the initiative during hunts,
and making decisions, but Mech could not identify the individual well enough
to know whether it was always the same Wolf. It did, however, seem to be a
male, and during the breeding season, a female sometimes led, with a male
close behind her.

Later, Jordan et al. (1967) recognized a particular male that led the Isle
Royale pack during travels for several winters. On the other hand, Peterson
(1977), observing the Wolves during the breeding season, believed that females
tended to lead the packs, as concluded by Pulliainen (1965: 236) who cited
anecdotal literature.

Haber (1977) considered a "beta male" to be the leader of the Savage Pack
in Denali Park. That animal tended to break trail, set the direction and pace
of travel and resting, and initiated and ended most of the hunts and rest
periods. However, this pack was highly unusual in that both the alpha male
and beta male remained with the pack until 8 or 9-years old. No one else has
ever reported two adult males remaining concurrently with a pack for even
4 years, including during a study of 13 packs for 4-9 years in the same area
(Mech et al. 1998). In two other packs, Haber believed that high-ranking males
generally led pack travels. During summer, leadership was less clear but Haber
believed it also tended to involve high-ranking males. None of these studies
provided quantitative behavioral data on leadership.